Panthera spelaea: Difference between revisions

Panthera spelaea; Temporal range: Middle-Late Pleistocene, 0.6–0.013 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Skeleton in Natural History Museum, Vienna
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	†P. spelaea
Binomial name
	†Panthera spelaea; Goldfuss, 1810
Subspecies
	Panthera spelaea vereshchagini (Baryshnikov & Boeskorov, 2001); Panthera spelaea spelaea Goldfuss, 1810;
	Red indicates the maximal range of Panthera spelaea, blue Panthera atrox, and green Panthera leo.
Synonyms
	Panthera leo spelaea M. Boule & L. De Villeneuve, 1927;

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Panthera spelaea, also known as the cave lion or steppe lion,^[1] is an extinct Panthera species that most likely evolved in Europe after the third Cromerian interglacial stage, less than 600,000 years ago. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion (Panthera leo) occurring in Africa and Asia,^[2] with the genetic divergence between the two species variously estimated between 1.9 million^[3]^[4] and 600,000 years ago.^[5] It is closely related and probably ancestral to the American lion (Panthera atrox).^[5] The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna. It became extinct about 13,000 years ago.^[6]

Taxonomy[edit]

Felis spelaea was the scientific name used by Georg August Goldfuss in 1810 for a fossil lion skull that was excavated in a cave in southern Germany.^[7] It possibly dates to the Würm glaciation.^[8]^[2]^[6]

Several authors regarded Panthera spelaea as a subspecies of the modern lion, and therefore as Panthera leo spelaea.^[9]^[10]^[11]^[2] One author considered the cave lion to be more closely related to the tiger based on a comparison of skull shapes, and proposed the scientific name Panthera tigris spelaea.^[12] Some authors regard the larger Middle Pleisto

Results from morphological studies showed that it is distinct in cranial and dental anatomy to justify the specific status of Panthera spelaea.^[13]^[14] Results of phylogenetic studies also support this assessment.^[15]^[3]^[4]

In 2001, the subspecies Panthera spelaea vereshchagini was proposed for seven specimens found in Siberia and Yukon, which have smaller skulls and teeth than the average P. spelaea.^[16] Before 2020, genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from P. spelaea from Europe and Alaska were indistinguishable, suggesting one large panmictic population.^[3]^[17] However, analysis of mitochondrial genome sequences from 31 cave lions showed that they fall into two monophyletic clades. One lived across western Europe and the other was restricted to Beringia during the Pleistocene. For this reason, the Beringian population is considered a distinct subspecies, P. s. vereshchagini.^[17]^[18]

Evolution[edit]

Lion-like pantherine felids first appeared in the Tanzanian Olduvai Gorge about 1.7 to 1.2 million years ago. These cats dispersed to Europe from East Africa in the first half of the Middle Pleistocene, giving rise to Panthera fossilis in Central Europe by 610,000 years ago.^[20] Different authors considered Panthera fossils as either a distinct species ancestral to P. spelaea,^[21] or as a subspecies of P. spelaea.^[22] Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.^[2]^[20]^[5]

The following cladogram shows the genetic relationship between P. spelaea and other pantherine cats.^[4]

Felis catus

Pantherinae

Neofelis nebulosa (clouded leopard)

Panthera

Panthera tigris (tiger)

Panthera onca (jaguar)

Panthera uncia (snow leopard)

Panthera pardus (leopard)

	Panthera leo leo (Asian, North and West African lions)

	Panthera leo melanochaita

(Southern and East African lions)

†Panthera spelaea

Specimens intermediate between P. fossilis and Late Pleistocene P. spelaea are referred to as the subspecies P. s. intermedia.^[23] The transition from P. fossilis to Late Pleistocene P. spelaea involved significant changes in skull and tooth morphology.^[24] Mitochondrial DNA sequence data from fossil lion remains show that the American lion represents a sister group of P. spelaea, and likely arose when an early P. spelaea population became isolated south of the Cordilleran Ice Sheet about 340,000 years ago.^[3]

Characteristics[edit]

Cave lions, Chamber of Felines, Lascaux caves

Carvings and cave paintings of cave lions, which were discovered in the Lascaux and Chauvet Caves in France, were dated to 15,000 to 17,000 years old.^[19]^[25] A drawing in the Chauvet cave depicts two cave lions walking together. The one in the foreground is slightly smaller than the one in the background, which has been drawn with a scrotum and without a mane.^[26] Such cave paintings suggest that male cave lions completely lacked manes, or at most had very small manes.^[6]

P. spelaea is also known from the Löwenmensch figurine found in Vogelherd cave in the Swabian Alb, southwest Germany, which dates to the Aurignacian culture. These archaeological artifacts indicate that it may have been featured in Paleolithic religious rituals.^[27]^[28]

P. spelaea was thought to have been one of the largest lion species. The skeleton of an adult male found in 1985 near Siegsdorf in Germany had a shoulder height of around 1.2 m (3 ft 11 in) and a head-body length of 2 m (6 ft 7 in) without the tail, similar in size to large modern lions.^[29] The size of this male was exceeded by other specimens, with another male reaching 2.5 m (8 ft 2 in) long without the tail.^{[citation needed]} Similarly, footprints attributed to a male cave lion measured 15 cm (6 in) across. The heaviest Panthera spelaea was estimated to weigh 339 kg (747 lb).^[30]

This shows that P. spelaea would have been up to or over 12% larger than modern lions, but still smaller than the earlier Panthera fossilis or the American lion (P. atrox). Cave paintings almost exclusively show hunting animals without a mane, suggesting that males were indeed maneless.^[31]^[32] P. spelaea had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.^[6] Like modern lions, females were smaller than males.^[33]

In 2016, hair found near the Maly Anyuy River was identified as cave lion hair through DNA analysis. Comparison with hair of a modern lion revealed that cave lion hair was probably similar in colour as that of the modern lion, though slightly lighter. In addition, the cave lion is thought to have had a very thick and dense undercoat comprising closed and compressed yellowish-to-white wavy downy hair with a smaller mass of darker-coloured guard hairs, possibly an adaptation to the Ice Age climate.^[34]

Distribution and habitat[edit]

P. spelaea formed a contiguous population from western Europe to the Yukon over the Bering land bridge, across the range of the mammoth steppe.^[6]^[35] It was widely distributed in the Iberian Peninsula,^[36] Italian Peninsula,^[37] Southeast Europe,^[38] Great Britain,^[6] Central Europe,^[39]^[40]^[41] the East European Plain,^[6] the Ural Mountains,^[42] and across most of northern Eurasia into Alaska and Yukon.^[6] The cave lion had a wide elevation range as well; in Slovakia, skull, femur and pelvis remains were excavated in ten Karst caves in hilly and montane areas at elevations from 240 to 1,133 m (787 to 3,717 ft).^[43]

The oldest known fossils dated with radiocarbon dating were excavated in northeastern Russia and were radiocarbon dated at 62,400 years old.^[6] Phalanx bones excavated in Spain's La Garma cave complex were radiocarbon dated to 14,300–14,000 years old.^[44] The youngest known fossils are dated 11,925 years old and originated near Fairbanks, Alaska.^[6] In Western Europe, however, P. spelaea may have survived into the Early Holocene.^[45]

In 2008, a well-preserved mature cave lion specimen was unearthed near the Maly Anyuy River in Chukotka Autonomous Okrug, which still retained some clumps of hair.^[46] In the Sakha Republic's Khayrgas Cave, bones of P. spelaea were found together with remains of humans, wolf, reindeer, Pleistocene horse and fish in a layer dated 13,200–21,500 years old.^[47]

The cave lion probably inhabited predominantly open habitats such as steppe and grasslands although it would have also have occurred in open woodlands as well.^[6] It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.^[48] While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,^[49] such as in Europe during the Last Interglacial/Eemian.^[50]

Discoveries[edit]

In 2015, two frozen cave lion cubs, estimated to be between 25,000 and 55,000 years old, were discovered close to the Uyandina River in Yakutia, Siberia in permafrost.^[51]^[52]^[53] Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.^[54]

In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River (Russian: Тирехтях), a tributary of the Indigirka River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.^[55] In 2018, another preserved carcass of a cub was found in a location 15 m (50 ft) away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.^[56] However, carbon dating showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.^[18] Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the Ice age.^[57]

Paleobiology[edit]

P. spelaea inhabited colder environments such as mammoth steppe and boreal forest. It was one of the keystone species of the mammoth steppe, being one of the main apex predators alongside the gray wolf, cave hyena and brown bear.^[59] Large amounts of bones belonging to P. spelaea were excavated in caves, where bones of cave hyena, cave bear and Paleolithic artefacts were also found.^[60]^[61] Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.^[62] It is unclear whether P. spelaea was social like the modern lion; some evidence indicates that it may have been solitary.^[59]

Isotopic analyses of bone collagen samples extracted from fossils indicate that cave bear cubs, reindeer and other cervids (deer) were prominent in the diet of cave lions. Later cave lions seem to have preyed foremost on reindeer, up to the brink of local extinction or extirpation of both species.^[59]^[63] Other possible prey species were giant deer, red deer, wild horse, muskox, aurochs, wisent, steppe bison, young woolly rhino, and young woolly mammoth. It likely competed for prey with the European Ice Age leopard, cave hyenas, brown bears and grey wolves in Eurasia,^[64] along with short-faced bears, Homotherium, and Beringian wolves in Beringia.^[59]

Extinction[edit]

The extinction of the Eurasian cave lion was multifaceted, being attributable to a combination of human activity, the decline of the mammoth steppe, and changes in the herbivore guild.^[23] There is direct evidence of human hunting of cave lions and exploitation of their pelts for ritualistic purposes during the Magdalenian, which was likely an important factor contributing to their ultimate demise.^[44]

References[edit]

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^ Kuzmin, Y. V.; Kosintsev, P. A.; Stepanov, A. D.; Boeskorov, G. G.; Cruz, R. J. (2017). "Chronology and faunal remains of the Khayrgas Cave (Eastern Siberia, Russia)" (PDF). Radiocarbon. 59 (2): 575−582. Bibcode:2017Radcb..59..575K. doi:10.1017/RDC.2016.39. S2CID 133453976. Archived from the original (PDF) on 20 February 2020.
^ Sabol, M.; Gullár, J.; Horvát, J. (2018). "Montane record of the late Pleistocene Panthera spelaea (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia)". Journal of Vertebrate Paleontology. 38 (3): e1467921. Bibcode:2018JVPal..38E7921S. doi:10.1080/02724634.2018.1467921.
^ Álvarez-Lao, Diego J.; Álvarez-Vena, Adrián; Ballesteros, Daniel; García, Nuria; Laplana, César (February 2020). "A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3)". Quaternary Science Reviews. 229: 106123. doi:10.1016/j.quascirev.2019.106123. hdl:10651/54312.
^ Diedrich, Cajus G. (June 2011). "A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany". Historical Biology. 23 (2–3): 195–217. doi:10.1080/08912963.2010.507814. ISSN 0891-2963.
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^ "Whiskers still bristling after more than 12,000 years in the Siberian cold". The Siberian Times. 2015. Archived from the original on 16 December 2016. Retrieved 25 November 2016.
^ "Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes". The Siberian Times. 2017. Archived from the original on 12 November 2017. Retrieved 12 November 2017.
^ Gertcyk, O. (2018). "Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost". The Siberian Times. Archived from the original on 14 September 2018. Retrieved 16 September 2018.
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External links[edit]

Prehistoric cats and prehistoric cat-like creatures, from the Messybeast Cat Resource Archive
The mammoth and the flood, volume 5, chapter 1, by Hans Krause.
Hoyle and cavetigers Archived 13 January 2016 at the Wayback Machine, from the Dinosaur Mailing List. (Groiss)
Photo-reconstruction of Panthera spelaea, by paleoartist Roman Uchytel

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[62] Diedrich, Cajus G. (1 July 2011). "Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe". Quaternary Science Reviews. 30 (15): 1883–1906. Bibcode:2011QSRv...30.1883D. doi:10.1016/j.quascirev.2011.03.006. ISSN 0277-3791. Archived from the original on 18 May 2024. Retrieved 4 February 2024 – via Elsevier Science Direct.

[Bocherens-63] Bocherens, H.; Drucker, D. G.; Bonjean, D.; Bridault, A.; Conard, N. J.; Cupillard, C.; Germonpré, M.; Höneisen, M.; Münzel, S. C.; Napierala, H. & Patou-Mathis, M. (2011). "Isotopic evidence for dietary ecology of cave lion (Panthera spelaea) in North-Western Europe: prey choice, competition and implications for extinction" (PDF). Quaternary International. 245 (2): 249–261. Bibcode:2011QuInt.245..249B. doi:10.1016/j.quaint.2011.02.023. S2CID 129706533. Archived (PDF) from the original on 11 May 2020. Retrieved 18 September 2020.

[Diedrich_2013-64] Diedrich, C. G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.

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@@ Line 2: / Line 2: @@
 {{Use dmy dates|date=May 2017}}
 {{Speciesbox
-| fossil_range = [[Late Pleistocene]], {{Fossil range|0.6|0.013}}
+| fossil_range = [[Chibanian|Middle]]-[[Late Pleistocene]], {{Fossil range|0.6|0.013}}
 | image = Panthera leo spelaea in Vienna.jpg
 | image_caption = [[Skeleton]] in [[Natural History Museum, Vienna]]
@@ Line 12: / Line 12: @@
 | range_map_caption = Red indicates the maximal range of ''Panthera spelaea'', blue ''[[American lion|Panthera atrox]]'', and green ''[[Lion|Panthera leo]]''.
 | synonyms = *''Panthera leo spelaea'' <small>M. Boule & L. De Villeneuve, 1927</small>
-*''Uncia spelaea''
 | subdivision_ranks = Subspecies
+| subdivision = *''Panthera spelaea vereshchagini'' <small>(Baryshnikov & Boeskorov, 2001)</small>
-| subdivision =
-*''Panthera spelaea vereshchagini'' <small>(Baryshnikov & Boeskorov, 2001)</small>
 *''Panthera spelaea spelaea'' <small>Goldfuss, 1810</small>
 }}
-'''''Panthera spelaea''''', also known as the '''Eurasian cave lion''', '''European cave lion''' or '''steppe lion''',<ref name="diedrich2014">{{cite journal |last=Diedrich |first=C. G. |year= 2014 |title=Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey |journal=[[Paleontology Journal]] |volume=2014 |pages=1–34 |doi=10.1155/2014/106203 |doi-access=free}}</ref> is an [[extinct]] ''[[Panthera]]'' species that most likely evolved in [[Europe]] after the third [[Cromerian interglacial]] stage, less than 600,000 years ago. [[Phylogenetics|Phylogenetic]] analysis of [[fossil]] [[bone]] samples revealed that it was highly distinct and genetically isolated from the modern [[lion]] (''Panthera leo'') occurring in Africa and Asia.<ref name="Burger">{{cite journal |last1=Burger |first1=J. |last2=Rosendahl |first2=W. |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last10=Alt |first10=K. W. |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |volume=30 |issue=3 |pages=841–849 |year=2004 |doi=10.1016/j.ympev.2003.07.020 |pmid=15012963}}</ref> The species ranged from Western Europe to eastern [[Beringia]] in North America, and was a prominent member of the [[mammoth steppe]] fauna.
+'''''Panthera spelaea''''', also known as the '''cave lion''' or '''steppe lion''',<ref name="diedrich2014">{{cite journal |last=Diedrich |first=C. G. |year= 2014 |title=Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey |journal=[[Paleontology Journal]] |volume=2014 |pages=1–34 |doi=10.1155/2014/106203 |doi-access=free}}</ref> is an [[extinct]] ''[[Panthera]]'' species that most likely evolved in [[Europe]] after the third [[Cromerian interglacial]] stage, less than 600,000 years ago. Genetic analysis of [[ancient DNA]] has revealed that while closely related, it was a distinct species genetically isolated from the modern [[lion]] (''Panthera leo'') occurring in Africa and Asia,<ref name="Burger">{{cite journal |last1=Burger |first1=J. |last2=Rosendahl |first2=W. |last3=Loreille |first3=O. |last4=Hemmer |first4=H. |last5=Eriksson |first5=T. |last6=Götherström |first6=A. |last7=Hiller |first7=J. |last8=Collins |first8=M. J. |last9=Wess |first9=T. |last10=Alt |first10=K. W. |title=Molecular phylogeny of the extinct cave lion ''Panthera leo spelaea'' |journal=[[Molecular Phylogenetics and Evolution]] |volume=30 |issue=3 |pages=841–849 |year=2004 |doi=10.1016/j.ympev.2003.07.020 |pmid=15012963}}</ref> with the [[genetic divergence]] between the two species variously estimated between 1.9 million<ref name="Barnett2009">{{cite journal |author1=Barnett, R. |author2=Shapiro, B. |author3=Barnes, I. A. N. |author4=Ho, S. Y. |author5=Burger, J. |author6=Yamaguchi, N. |author7=Higham, T. F. |author8=Wheeler, H. |author9=Rosendahl, W. |author10=Sher, A. V. |author11=Sotnikova, M. |name-list-style=amp |year=2009 |title=Phylogeography of lions (''Panthera leo ssp.'') reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf |journal=[[Molecular Ecology]] |volume=18 |issue=8 |pages=1668–1677 |doi=10.1111/j.1365-294X.2009.04134.x |pmid=19302360 |bibcode=2009MolEc..18.1668B |s2cid=46716748 |archive-url=https://web.archive.org/web/20120121153712/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf |archive-date=21 January 2012}}</ref><ref name="Barnett2016">{{cite journal |last1=Barnett |first1=R. |last2=Mendoza |first2=M. L. Z. |last3=Soares |first3=A. E. R. |last4=Ho |first4=S. Y. W. |last5=Zazula |first5=G. |last6=Yamaguchi |first6=N. |last7=Shapiro |first7=B. |last8=Kirillova |first8=I. V. |last9=Larson |first9=G. |last10=Gilbert |first10=M. T. P. |year=2016 |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), Resolve its Position within the ''Panthera'' Cats |journal=Open Quaternary |volume=2 |page=4 |doi=10.5334/oq.24 |doi-access=free|hdl=10576/22920 |hdl-access=free }}</ref> and 600,000 years ago.<ref name="Manuel_al2020">{{cite journal |author1=Manuel, M. d. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |name-list-style=amp |year=2020 |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |bibcode=2020PNAS..11710927D |doi=10.1073/pnas.1919423117 |pmc=7245068 |pmid=32366643 |doi-access=free |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T. |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P.}}</ref> It is closely related and probably ancestral to the [[American lion]] (''Panthera atrox'').<ref name="Manuel_al2020" />  The species ranged from Western Europe to eastern [[Beringia]] in North America, and was a prominent member of the [[mammoth steppe]] fauna. It became extinct about 13,000 years ago.<ref name="Stuart2011">{{cite journal |author1=Stuart, A. J. |author2=Lister, A. M. |name-list-style=amp |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=[[Quaternary Science Reviews]] |volume=30 |issue=17 |pages=2329–2340 |bibcode=2011QSRv...30.2329S |doi=10.1016/j.quascirev.2010.04.023}}</ref>
-Analysis of [[Morphology (biology)|morphological]] differences and [[mitochondrial]] data support the [[Taxonomy (biology)|taxonomic]] recognition of ''Panthera spelaea'' as a distinct [[species]] that [[Genetic divergence|genetically diverged]] from the lion about {{mya|1.9}}.<ref name="Barnett2009">{{cite journal |author1=Barnett, R. |name-list-style=amp |author2=Shapiro, B. |author3=Barnes, I. A. N. |author4=Ho, S. Y. |author5=Burger, J. |author6=Yamaguchi, N. |author7=Higham, T. F. |author8=Wheeler, H. |author9=Rosendahl, W. |author10=Sher, A. V. |author11=Sotnikova, M. |year=2009 |title=Phylogeography of lions (''Panthera leo ssp.'') reveals three distinct taxa and a late Pleistocene reduction in genetic diversity |journal=[[Molecular Ecology]] |volume=18 |issue=8 |pages=1668–1677 |pmid=19302360 |doi=10.1111/j.1365-294X.2009.04134.x |s2cid=46716748 |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf  |archive-url=https://web.archive.org/web/20120121153712/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Barnett%20et%20al.%202009.pdf |archive-date=21 January 2012 }}</ref><ref name=Barnett2016>{{cite journal |last1=Barnett |first1=R. |last2=Mendoza |first2=M. L. Z. |last3=Soares |first3=A. E. R. |last4= Ho|first4=S. Y. W. |last5=Zazula |first5=G. |last6=Yamaguchi |first6=N. |last7=Shapiro |first7=B. |last8=Kirillova |first8=I. V. |last9=Larson |first9=G. |last10=Gilbert |first10=M. T. P. |title=Mitogenomics of the Extinct Cave Lion, ''Panthera spelaea'' (Goldfuss, 1810), Resolve its Position within the ''Panthera'' Cats |journal=Open Quaternary |volume=2 |year=2016 |page=4 |doi=10.5334/oq.24 |doi-access=free}}</ref> [[Nuclear genome|Nuclear genomic]] evidence shows a more recent split approximately 500,000 years ago, with no subsequent interbreeding with the ancestors of the modern lion. It is closely related and probably ancestral to the [[American lion]] (''Panthera atrox'').<ref name=Manuel_al2020>{{cite journal |name-list-style=amp |author1=Manuel, M. d. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T. |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O'Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P. |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |volume=117 |issue=20 |pages=10927–10934 |doi=10.1073/pnas.1919423117 |year=2020 |pmid=32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref> It became extinct about 13,000 years ago.<ref name="Stuart2011">{{cite journal |author1=Stuart, A. J. |name-list-style=amp |author2=Lister, A. M. |year=2011 |title=Extinction chronology of the cave lion ''Panthera spelaea'' |journal=Quaternary Science Reviews |volume=30 |issue=17 |pages=2329–2340 |doi=10.1016/j.quascirev.2010.04.023 |bibcode=2011QSRv...30.2329S}}</ref>
 ==Taxonomy==
@@ Line 28: / Line 24: @@
 Several authors regarded ''Panthera spelaea'' as a [[subspecies]] of the modern [[lion]], and therefore as ''Panthera leo spelaea''.<ref>{{cite book |author=Kurtén, B. |date=1968 |title=Pleistocene Mammals of Europe |publisher=Weidenfeld and Nicolson |location=London}}</ref><ref name=Hemmer>{{cite journal |author=Hemmer, H. |year=1974 |title=Untersuchungen zur Stammesgeschichte der Pantherkatzen (''Pantherinae'') Teil 3. Zur Artgeschichte des Löwen ''Panthera (Panthera) leo'' (Linnaeus, 1758) |journal=Veröffentlichungen der Zoologischen Staatssammlung 17 |pages=167–280 |url=https://archive.org/stream/verfentlichungen171974zool#page/178/mode/2up}}</ref><ref>{{cite journal |author=Turner, A. |year=1984 |title=Dental sex dimorphism in European lions (''Panthera leo'' L.) of the Upper Pleistocene: palaeoecological and palaeoethological implications |journal=Annales Zoologici Fennici |volume=21 |pages=1–8}}</ref><ref name=Burger/>
-One author considered the cave lion to be more closely related to the [[tiger]] based on a comparison of [[skull]] shapes, and proposed the scientific name ''Panthera tigris spelaea''.<ref>{{cite journal |author=Groiss, J. Th. |year=1996 |title=Der Höhlentiger ''Panthera tigris spelaea'' (Goldfuss) |journal=[[Neues Jahrbuch für Geologie und Paläontologie]] |issue=7 |pages=399–414 |doi=10.1127/njgpm/1996/1996/399}}</ref>
+One author considered the cave lion to be more closely related to the [[tiger]] based on a comparison of [[skull]] shapes, and proposed the scientific name ''Panthera tigris spelaea''.<ref>{{cite journal |author=Groiss, J. Th. |year=1996 |title=Der Höhlentiger ''Panthera tigris spelaea'' (Goldfuss) |journal=[[Neues Jahrbuch für Geologie und Paläontologie]] |issue=7 |pages=399–414 |doi=10.1127/njgpm/1996/1996/399}}</ref> Some authors regard the larger Middle Pleisto
-Results from morphological studies showed that it is distinct in cranial and dental [[anatomy]] to justify the [[Species|specific]] status of ''Panthera spelaea''.<ref>{{cite book |author=Spassov, N. |author2=Iliev, N. |name-list-style=amp |year=1994 |title=Proceedings of the International Symposium VI |location=Thracia Pontica |chapter=Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast) |pages=287–314}}</ref><ref>{{cite journal |author=Sotnikova, M. |author2=Nikolskiy, P. |year=2006 |title=Systematic position of the cave lion ''Panthera spelaea'' (Goldfuss) based on cranial and dental characters |journal=Quaternary International |volume=142-143 |pages=218–228 |doi=10.1016/j.quaint.2005.03.019 |bibcode=2006QuInt.142..218S|url=http://doc.rero.ch/record/13484/files/PAL_E260.pdf }}</ref> Results of [[Phylogenetics|phylogenetic]] studies also support this assessment.<ref>{{cite journal |last1=Christiansen |first1=P. |year=2008 |title=Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters |journal=[[Cladistics (journal)|Cladistics]] |volume=24 |issue=6 |pages=977–992 |doi=10.1111/j.1096-0031.2008.00226.x |pmid=34892880 |s2cid=84497516}}</ref><ref name=Barnett2009 /><ref name=Barnett2016/>
+Results from morphological studies showed that it is distinct in cranial and dental [[anatomy]] to justify the [[Species|specific]] status of ''Panthera spelaea''.<ref>{{cite book |author=Spassov, N. |author2=Iliev, N. |name-list-style=amp |year=1994 |title=Proceedings of the International Symposium VI |location=Thracia Pontica |chapter=Animal remains from the submerged Late Eneolithic – early Bronze Age settlements in Sozopol (South Bulgarian Black Sea Coast) |pages=287–314}}</ref><ref>{{cite journal |author=Sotnikova, M. |author2=Nikolskiy, P. |year=2006 |title=Systematic position of the cave lion ''Panthera spelaea'' (Goldfuss) based on cranial and dental characters |journal=Quaternary International |volume=142-143 |pages=218–228 |doi=10.1016/j.quaint.2005.03.019 |bibcode=2006QuInt.142..218S |url=http://doc.rero.ch/record/13484/files/PAL_E260.pdf |access-date=22 January 2023 |archive-date=5 February 2023 |archive-url=https://web.archive.org/web/20230205061457/https://doc.rero.ch/record/13484/files/PAL_E260.pdf |url-status=live }}</ref> Results of [[Phylogenetics|phylogenetic]] studies also support this assessment.<ref>{{cite journal |last1=Christiansen |first1=P. |year=2008 |title=Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters |journal=[[Cladistics (journal)|Cladistics]] |volume=24 |issue=6 |pages=977–992 |doi=10.1111/j.1096-0031.2008.00226.x |pmid=34892880 |s2cid=84497516|doi-access=free }}</ref><ref name=Barnett2009 /><ref name=Barnett2016/>
-In 2001, the subspecies ''Panthera spelaea vereshchagini'' was proposed for seven specimens found in [[Siberia]] and [[Yukon]], which have smaller skulls and teeth than the average ''P. spelaea''.<ref>{{cite journal |author1=Baryshnikov, G. F. |author2=Boeskorov, G. |name-list-style=amp |year=2001 |title=The Pleistocene cave lion, ''Panthera spelaea'' (Carnivora, Felidae) from Yakutia, Russia |journal=Cranium |volume=18 |issue=1 |pages=7–24 |url=http://natuurtijdschriften.nl/download?type=document;docid=523468}}</ref> Before 2020, genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from ''P. spelaea'' from Europe and Alaska were indistinguishable, suggesting one large [[panmictic]] population.<ref name=Barnett2009/><ref name=Stanton2020>{{cite journal |author1=Stanton, D.W. |name-list-style=amp |author2=Alberti, F. |author3=Plotnikov, V. |author4=Androsov, S. |author5=Grigoriev, S. |author6=Fedorov, S. |author7=Kosintsev, P. |author8=Nagel, D. |author9=Vartanyan, S. |author10=Barnes, I. |author11=Barnett, R. |journal=Scientific Reports |volume=10 |page=12621 |year=2020 |title=Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion |issue=1 |doi=10.1038/s41598-020-69474-1 |pmid=32724178 |pmc=7387438 |bibcode=2020NatSR..1012621S |doi-access=free}}</ref> However, analysis of [[mitochondrial genome]] sequences from 31 cave lions showed that they fall into two [[Monophyletic group|monophyletic]] [[clade]]s. One lived across western Europe and the other was restricted to Beringia during the [[Pleistocene]]. For this reason, the Beringian population is considered a distinct subspecies, ''P. s. vereshchagini''.<ref name=Stanton2020 /><ref name="Boeskorov_al.2021">{{cite journal |last1=Boeskorov |first1=G. G. |last2=Plotnikov |first2=V. V. |last3=Protopopov |first3=A. V. |last4=Baryshnikov |first4=G. F. |last5=Fosse |first5=P. |last6=Dalén |first6=L. |last7=David |first7=S. W. G. |last8=Pavlov |first8=I. S. |last9=Naoki |first9=S. |last10=Alexey |first10=N. T. |name-list-style=amp |title=The preliminary analysis of Cave Lion cubs ''Panthera spelaea'' (Goldfuss, 1810) from the permafrost of Siberia |journal=[[Quaternary (journal)|Quaternary]] |volume=4 |issue=3 |page=24 |doi=10.3390/quat4030024 |doi-access=free |date=2021}}</ref>
+In 2001, the subspecies ''Panthera spelaea vereshchagini'' was proposed for seven specimens found in [[Siberia]] and [[Yukon]], which have smaller skulls and teeth than the average ''P. spelaea''.<ref>{{cite journal |author1=Baryshnikov, G. F. |author2=Boeskorov, G. |name-list-style=amp |year=2001 |title=The Pleistocene cave lion, ''Panthera spelaea'' (Carnivora, Felidae) from Yakutia, Russia |journal=Cranium |volume=18 |issue=1 |pages=7–24 |url=http://natuurtijdschriften.nl/download?type=document;docid=523468 |access-date=31 December 2017 |archive-date=1 January 2018 |archive-url=https://web.archive.org/web/20180101030312/http://natuurtijdschriften.nl/download?type=document;docid=523468 |url-status=live }}</ref> Before 2020, genetic analysis using ancient DNA provided no evidence for their distinct subspecific status; DNA signatures from ''P. spelaea'' from Europe and Alaska were indistinguishable, suggesting one large [[panmictic]] population.<ref name=Barnett2009/><ref name=Stanton2020>{{cite journal |author1=Stanton, D.W. |name-list-style=amp |author2=Alberti, F. |author3=Plotnikov, V. |author4=Androsov, S. |author5=Grigoriev, S. |author6=Fedorov, S. |author7=Kosintsev, P. |author8=Nagel, D. |author9=Vartanyan, S. |author10=Barnes, I. |author11=Barnett, R. |journal=Scientific Reports |volume=10 |page=12621 |year=2020 |title=Early Pleistocene origin and extensive intra-species diversity of the extinct cave lion |issue=1 |doi=10.1038/s41598-020-69474-1 |pmid=32724178 |pmc=7387438 |bibcode=2020NatSR..1012621S |doi-access=free}}</ref> However, analysis of [[mitochondrial genome]] sequences from 31 cave lions showed that they fall into two [[Monophyletic group|monophyletic]] [[clade]]s. One lived across western Europe and the other was restricted to Beringia during the [[Pleistocene]]. For this reason, the Beringian population is considered a distinct subspecies, ''P. s. vereshchagini''.<ref name=Stanton2020 /><ref name="Boeskorov_al.2021">{{cite journal |last1=Boeskorov |first1=G. G. |last2=Plotnikov |first2=V. V. |last3=Protopopov |first3=A. V. |last4=Baryshnikov |first4=G. F. |last5=Fosse |first5=P. |last6=Dalén |first6=L. |last7=David |first7=S. W. G. |last8=Pavlov |first8=I. S. |last9=Naoki |first9=S. |last10=Alexey |first10=N. T. |name-list-style=amp |title=The preliminary analysis of Cave Lion cubs ''Panthera spelaea'' (Goldfuss, 1810) from the permafrost of Siberia |journal=[[Quaternary (journal)|Quaternary]] |volume=4 |issue=3 |page=24 |doi=10.3390/quat4030024 |doi-access=free |date=2021}}</ref>
 ===Evolution===
 [[File:18 PanneauDesLions(PartieDroite)BisonsPoursuivisParDesLions.jpg|thumb|Cave lions and [[bison]] depicted in the [[Chauvet Cave]], [[France]]<ref name=Chauvet1996>{{cite book |author=Chauvet, J.-M. |author2=Brunel, D. E. |author3=Hillaire, C. |name-list-style=amp |year=1996 |title=Dawn of Art: The Chauvet Cave. The oldest known paintings in the world |url=https://archive.org/details/dawnofartchauvet0000chau |url-access=registration |location=New York |publisher=Harry N. Abrams}}</ref>]]
-Lion-like [[Panthera|pantherine]] felids first appeared in the [[Tanzania]]n [[Olduvai Gorge]] about {{mya|1.7|1.2}}. These cats dispersed to Europe from [[East Africa]] in the first half of the  Middle  Pleistocene, giving rise to ''[[Panthera leo fossilis|P. fossilis]]'' in [[Central Europe]] by 610,000 years ago.<ref name=Sotnikova2014>{{cite journal |author1=Sotnikova, M.V. |author2=Foronova, I.V. |name-list-style=amp |year=2014 |title=First Asian record of ''Panthera (Leo) fossilis'' (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia |journal=Integrative Zoology |volume=9  |issue=4 |pages=517–530|doi=10.1111/1749-4877.12082 |pmid=24382145}}</ref> ''Panthera spelaea'' evolved from ''P. fossilis'' about 460,000 years ago in central Europe during the late [[Saalian glaciation]] or early [[Eemian]] and would have been common throughout [[Eurasia]] from 450,000 to 14,000 years ago. Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.<ref name=Burger/><ref name=Sotnikova2014/><ref name=Manuel_al2020/>
+Lion-like [[Panthera|pantherine]] felids first appeared in the [[Tanzania]]n [[Olduvai Gorge]] about {{mya|1.7|1.2}}. These cats dispersed to Europe from [[East Africa]] in the first half of the Middle Pleistocene, giving rise to ''[[Panthera fossilis]]'' in [[Central Europe]] by 610,000 years ago.<ref name=Sotnikova2014>{{cite journal |author1=Sotnikova, M.V. |author2=Foronova, I.V. |name-list-style=amp |year=2014 |title=First Asian record of ''Panthera (Leo) fossilis'' (Mammalia, Carnivora, Felidae) in the Early Pleistocene of Western Siberia, Russia |journal=Integrative Zoology |volume=9  |issue=4 |pages=517–530|doi=10.1111/1749-4877.12082 |pmid=24382145}}</ref> Different authors considered ''Panthera fossils'' as either a distinct species ancestral to ''P. spelaea'',<ref name=":02">{{Cite journal |last1=Sabo |first1=Martin |last2=Tomašových |first2=Adam |last3=Gullár |first3=Juraj |date=August 2022 |title=Geographic and temporal variability in Pleistocene lion-like felids: Implications for their evolution and taxonomy |url=https://palaeo-electronica.org/content/2022/3681-lion-cranial-variability |journal=Palaeontologia Electronica |language=English |volume=25 |issue=2 |pages=1–27 |doi=10.26879/1175 |issn=1094-8074 |s2cid=251855356 |doi-access=free}}</ref> or as a subspecies of ''P. spelaea''.<ref name=":1">{{Cite journal |last=Marciszak |first=Adrian |last2=Ivanoff |first2=Dmitry V. |last3=Semenov |first3=Yuriy A. |last4=Talamo |first4=Sahra |last5=Ridush |first5=Bogdan |last6=Stupak |first6=Alina |last7=Yanish |first7=Yevheniia |last8=Kovalchuk |first8=Oleksandr |date=March 2023 |title=The Quaternary lions of Ukraine and a trend of decreasing size in Panthera spelaea |url=https://link.springer.com/10.1007/s10914-022-09635-3 |journal=Journal of Mammalian Evolution |language=en |volume=30 |issue=1 |pages=109–135 |doi=10.1007/s10914-022-09635-3 |issn=1064-7554|doi-access=free }}</ref> Recent nuclear genomic evidence suggest that interbreeding between modern lions and all Eurasian fossil lions took place up until 500,000 years ago, but by 470,000 years ago, no subsequent interbreeding between the two lineages occurred.<ref name=Burger/><ref name=Sotnikova2014/><ref name=Manuel_al2020/>
+The following cladogram shows the genetic relationship between ''P. spelaea'' and other pantherine cats.<ref name="Barnett2016" />{{clade| style=font-size:100%; line-height:100%
-''P. spelaea'' bone fragments excavated in [[Poland]] were radiocarbon dated to between the early and late [[Weichselian glaciation]], and are between 109,000 and 57,000 years old.<ref name=Marciszak2010>{{cite journal |author1=Marciszak, A. |name-list-style=amp |author2=Stefaniak, K. |year=2010 |title=Two forms of cave lion: Middle Pleistocene ''Panthera spelaea fossilis'' Reichenau, 1906 and Upper Pleistocene ''Panthera spelaea spelaea'' Goldfuss, 1810 from the Bisnik Cave, Poland |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=258 |issue=3 |pages=339–351 |doi=10.1127/0077-7749/2010/0117}}</ref> In Eurasia, it became extinct between 14,900 and 14,100 years ago, and survived in [[Beringia]] until 13,800 to 13,300 years ago as the Weichselian glaciation receded.<ref name=Stuart2011/><ref name=Manuel_al2020/> [[Mitochondrial DNA]] sequence data from fossil lion remains show that the [[American lion]] represents a [[sister group]] of ''P. spelaea'', and likely arose when an early ''P. spelaea'' population became isolated south of the [[Cordilleran Ice Sheet]] about 340,000 years ago.<ref name=Barnett2009/>
-The following cladogram shows the genetic relationship between ''P. spelaea'' and other pantherine cats.<ref name=Barnett2016/>
-{{clade| style=font-size:100%; line-height:100%
 |1={{clade
  |1=''[[Cat|Felis catus]]''[[File:Felis caligata - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam -(white background).jpg|50 px]]
+|label2=[[Pantherinae]]
  |2={{clade
-  |1=''[[Neofelis nebulosa]]''[[File:Studienblatt Felis macroscelis Nebelparder (white background).jpg|70 px]]
+  |1=''[[Neofelis nebulosa]]''[[File:Studienblatt Felis macroscelis Nebelparder (white background).jpg|70 px]] (clouded leopard)
+|label2=''[[Panthera]]''
   |2={{clade
+   |1=''[[Panthera tigris]]''[[File:Stamp-russia2014-save-russian-cats-(tiger).png|90 px]] (tiger)
-   |1={{clade
-    |1=''[[Panthera tigris sondaica]]'' [[File:Javan-Tiger-Panthera-tigris-sondaica (white background).jpg|70 px]]
-    |2=''[[Panthera tigris tigris]]''[[File:Stamp-russia2014-save-russian-cats-(tiger).png|90 px]] }}
    |2={{clade
-    |1=''[[Panthera onca]]''[[File:Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]
+    |1=''[[Panthera onca]]''[[File:Felis onca - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]] (jaguar)
     |2={{clade
-     |1=''[[Panthera uncia]]''[[File:Stamp-russia2014-save-russian-cats-(snow leopard).png|70 px]]
+     |1=''[[Panthera uncia]]''[[File:Stamp-russia2014-save-russian-cats-(snow leopard).png|70 px]] (snow leopard)
      |2={{clade
+      |1=''[[Panthera pardus]]''[[File:Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]] (leopard)
-      |1={{clade
-       |1=''[[Panthera pardus pardus]]''[[File:Felis pardus - 1818-1842 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).jpg|70 px]]
-       |2=''[[Panthera pardus orientalis]]''[[File:Stamp-russia2014-save-russian-cats-(leopard).png|70 px]]}}
       |2={{clade
        |1={{clade
-        |1=''[[Panthera leo leo]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo leo ♂.png|70 px]]</span>
+        |1=''[[Panthera leo leo]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo leo ♂.png|70 px]]</span> (Asian, North and West African lions)
-        |2=''[[Panthera leo melanochaita]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo melanochaita ♂.png|70 px]]</span>}}
+        |2=''[[Panthera leo melanochaita]]''<span style="{{MirrorH}}">[[File:W. Gornig - P. leo melanochaita ♂.png|70 px]]</span>}} (Southern and East African lions)
+       |2={{extinct}}'''''Panthera spelaea'''''<span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea.png|90 px]]</span> }} }} }} }} }} }} }} }}Specimens intermediate between ''P. fossilis'' and Late Pleistocene ''P. spelaea'' are referred to as the subspecies ''P. s. intermedia''.<ref name=":0" /> The transition from ''P. fossilis'' to Late Pleistocene ''P. spelaea'' involved significant changes in skull and tooth morphology.<ref>{{Cite journal |last=Persico |first=Davide |date=June 2021 |title=First fossil record of cave lion (Panthera (Leo) spelaea intermedia) from alluvial deposits of the Po River in northern Italy |url=https://linkinghub.elsevier.com/retrieve/pii/S1040618221000902 |journal=Quaternary International |language=en |volume=586 |pages=14–23 |doi=10.1016/j.quaint.2021.02.029}}</ref> [[Mitochondrial DNA]] sequence data from fossil lion remains show that the [[American lion]] represents a [[sister group]] of ''P. spelaea'', and likely arose when an early ''P. spelaea'' population became isolated south of the [[Cordilleran Ice Sheet]] about 340,000 years ago.<ref name="Barnett2009" />
-       |2='''''Panthera spelaea'''''<span style="{{MirrorH}}">[[File:W. Gornig - P. spelaea spelaea.png|90 px]]</span> }} }} }} }} }} }} }} }}
 ==Characteristics==
@@ Line 69: / Line 61: @@
 ''P. spelaea'' is also known from the [[Löwenmensch figurine]] found in Vogelherd cave in the [[Swabian Alb]], southwest Germany, which dates to the [[Aurignacian]] culture. These [[Artifact (archaeology)|archaeological artifact]]s indicate that it may have been featured in Paleolithic religious [[ritual]]s.<ref>{{cite book |author1=Bahn, P. G. |author2=Vertut, J. |year=1997 |title=Journey Through the Ice Age |publisher=Weidenfeld & Nicolson |location=London |isbn=978-0-297-83588-2 |url=https://archive.org/details/journeythroughic00bahn |url-access=registration}}</ref><ref>{{cite book |author=Guthrie, R. D. |year=2005 |title=The Nature of Paleolithic Art |publisher=University of Chicago Press |location=Chicago}}</ref>
-''P. spelaea'' was thought to have been one of the largest lion species. The skeleton of an adult male found in 1985 near [[Siegsdorf]] in Germany had a shoulder height of around {{convert|1.2|m|ftin|abbr=on}} and a head-body length of {{convert|2|m|ftin|abbr=on}} without the tail, similar in size to large modern lions.<ref>{{cite journal|first1=S.T.|last1=Hussain|first2=H.|last2=Floss|year=2015|title=Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"|journal=Quartär|volume=65|pages=85–120|doi=10.7485/QU62_4}}</ref> The size of this male was exceeded by other specimens, with another male reaching {{convert|2.5|m|ftin|abbr=on}} long without the tail.{{cn|date=March 2023}} Similarly, footprints attributed to a male cave lion  measured  {{convert|15|cm|in|0|abbr=on}} across. The heaviest  ''Panthera spelaea'' was estimated to weigh {{convert|339|kg|lb|abbr=on}}.<ref>{{cite journal |last=Sherani |first=Shaheer |title=A new specimen-dependent method of estimating felid body mass |journal=PeerJ Preprints | year=2016 | page=16 |doi=10.7287/peerj.preprints.2327v2|url=https://peerj.com/preprints/2327.pdf |doi-access=free }}</ref>
+''P. spelaea'' was thought to have been one of the largest lion species. The skeleton of an adult male found in 1985 near [[Siegsdorf]] in Germany had a shoulder height of around {{convert|1.2|m|ftin|abbr=on}} and a head-body length of {{convert|2|m|ftin|abbr=on}} without the tail, similar in size to large modern lions.<ref>{{cite journal|first1=S.T.|last1=Hussain|first2=H.|last2=Floss|year=2015|title=Sharing the world with mammoths, cave lions and other beings: linking animal-human interactions and the Aurignacian "belief world"|journal=Quartär|volume=65|pages=85–120|doi=10.7485/QU62_4}}</ref> The size of this male was exceeded by other specimens, with another male reaching {{convert|2.5|m|ftin|abbr=on}} long without the tail.{{cn|date=March 2023}} Similarly, footprints attributed to a male cave lion  measured  {{convert|15|cm|in|0|abbr=on}} across. The heaviest  ''Panthera spelaea'' was estimated to weigh {{convert|339|kg|lb|abbr=on}}.<ref>{{cite journal |last=Sherani |first=Shaheer |title=A new specimen-dependent method of estimating felid body mass |journal=PeerJ Preprints |year=2016 |page=16 |doi=10.7287/peerj.preprints.2327v2 |url=https://peerj.com/preprints/2327.pdf |doi-access=free |access-date=15 March 2021 |archive-date=20 September 2022 |archive-url=https://web.archive.org/web/20220920173645/https://peerj.com/preprints/2327.pdf |url-status=dead }}</ref>
 This shows that ''P. spelaea''  would have been up to or over 12% larger than modern lions, but still smaller than the earlier ''Panthera fossilis'' or the [[American lion]] (''P. atrox''). [[Cave painting]]s almost exclusively show hunting animals without a mane, suggesting that males were indeed maneless.<ref name="Koenigswald02">{{Cite book |last=Koenigswald |first=W. v. |year=2002 |title=Lebendige Eiszeit: Klima und Tierwelt im Wandel |location=Stuttgart |publisher=Theiss |isbn=978-3-8062-1734-6|language=de}}</ref><ref>{{cite book |author=Koenigswald, W. v. |date=2002 |title=Lebendige Eiszeit|location=Darmstadt |publisher=Theiss-Wissenschaftliche Buchgemeinschaft |isbn=3-8062-1734-3}}</ref> ''P. spelaea'' had a relatively longer and narrower muzzle compared to that of the extant lion. Despite this, the two species do not exhibit major differences in morphology.<ref name="Stuart2011"/> Like modern lions, females were smaller than males.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Late Pleistocene ''Panthera leo spelaea'' (Goldfuss, 1810) skeletons from the Czech Republic (central Europe); their pathological cranial features and injuries resulting from intraspecific fights, conflicts with hyenas, and attacks on cave bears |journal=Bulletin of Geosciences |volume=86 |issue=4 |pages=817–840 |doi=10.3140/bull.geosci.1263 |doi-access=free }}</ref>
@@ Line 77: / Line 69: @@
 ==Distribution and habitat==
 [[File:Ice age fauna of northern Spain - Mauricio Antón.jpg|thumb|Cave lions and other Ice Age fauna in northern Spain, by [[Mauricio Antón]]]]
+''P. spelaea'' formed a contiguous population from western Europe to the [[Yukon]] over the [[Bering land bridge]], across the range of the [[mammoth steppe]].<ref name=Stuart2011/><ref>{{cite journal |author=Ersmark, E. |author2=Orlando, L. |author3=Sandoval-Castellanos, E. |author4=Barnes, I. |author5=Barnett, R. |author6=Stuart, A. |author7=Lister, A. |author8=Dalén, L. |year=2015 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |journal=[[Open Quaternary]] |volume=1 |issue=1 |pages=Art. 4 |doi=10.5334/oq.aa |doi-access=free}}</ref> It was widely distributed in the [[Iberian Peninsula]],<ref>{{Cite journal |last1=Álvarez-Lao |first1=D. J. |last2=Álvarez-Vena |first2=A. |last3=Ballesteros |first3=D. |last4=García |first4=N. |last5=Laplana |first5=C. |date=2020 |title=A cave lion (''Panthera spelaea'') skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |journal=[[Quaternary Science Reviews]] |volume=229 |page=106123 |bibcode=2020QSRv..22906123A |doi=10.1016/j.quascirev.2019.106123 |hdl-access=free |hdl=10651/54312}}</ref> [[Italian Peninsula]],<ref>Iannucci, Alessio, Mecozzi, Beniamino, Pineda, Antonio, Sardella, Raffaele, Carpentieri, Marco, Rabinovich, Rivka, and Moncel Marie-Helene. "Early Occurence of (''Panthera spelaea'') at the Middle Pleistocene Acheulean site of Notarchirico (MIS 16, Italy). ''Journal of Quarternary Sciences'' 39:3 (2024): 4.</ref> [[Southeast Europe]],<ref>{{Cite journal |last1=Smith |first1=Geoff M. |last2=Spasov |first2=Rosen |last3=Martisius |first3=Naomi L. |last4=Sinet-Mathiot |first4=Virginie |last5=Aldeias |first5=Vera |last6=Rezek |first6=Zeljko |last7=Ruebens |first7=Karen |last8=Pederzani |first8=Sarah |last9=McPherron |first9=Shannon P. |last10=Sirakova |first10=Svoboda |last11=Sirakov |first11=Nikolay |last12=Tsanova |first12=Tsenka |last13=Hublin |first13=Jean-Jacques |date=December 2021 |title=Subsistence behavior during the Initial Upper Paleolithic in Europe: Site use, dietary practice, and carnivore exploitation at Bacho Kiro Cave (Bulgaria) |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248421001263 |journal=[[Journal of Human Evolution]] |language=en |volume=161 |pages=103074 |doi=10.1016/j.jhevol.2021.103074 |pmid=34628301 |access-date=22 March 2024 |via=Elsevier Science Direct}}</ref> [[Great Britain]],<ref name="Stuart2011" /> [[Central Europe]],<ref>{{Cite journal |last1=Sabol |first1=Martin |last2=Döppes |first2=Doris |date=January 2020 |title=The first radiocarbon dating of a cave lion fossil from the Slovakian Western Carpathians |url=https://www.researchgate.net/publication/338937589 |journal=Acta Geologica Slovaca |volume=11 |issue=1 |pages=11–14 |via=ResearchGate}}</ref><ref>{{Cite journal |last1=Diedrich |first1=Cajus G. |last2=Rathgeber |first2=Thomas |date=12 October 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss 1810) skeleton remains of the Upper Rhine Valley (SW Germany) and contributions to their sexual dimorphism, taphonomy and habitus |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |journal=[[Historical Biology]] |language=en |volume=24 |issue=1 |pages=1–28 |doi=10.1080/08912963.2010.549943 |issn=0891-2963 |access-date=24 March 2024 |via=Taylor and Francis Online |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071934/https://www.tandfonline.com/doi/abs/10.1080/08912963.2010.549943 |url-status=live }}</ref><ref>{{Cite journal |last1=Marciszak |first1=Adrian |last2=Schouwenburg |first2=Charles |last3=Gornig |first3=Wiktoria |last4=Lipecki |first4=Grzegorz |last5=Mackiewicz |first5=Paweł |date=1 November 2019 |title=Morphometric comparison of Panthera spelaea (Goldfuss, 1810) from Poland with the lion remains from Eurasia over the last 700 ka |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |journal=[[Quaternary Science Reviews]] |language=en |volume=223 |pages=105950 |doi=10.1016/j.quascirev.2019.105950 |bibcode=2019QSRv..22305950M |access-date=24 March 2024 |via=Elsevier Science Direct |archive-date=29 March 2023 |archive-url=https://web.archive.org/web/20230329081534/https://linkinghub.elsevier.com/retrieve/pii/S0277379119303907 |url-status=live }}</ref> the [[East European Plain]],<ref name="Stuart2011" /> the [[Ural Mountains]],<ref>{{Cite journal |last1=Gimranov |first1=Dmitry |last2=Kosintsev |first2=Pavel |date=30 April 2020 |title=Quaternary large mammals from the Imanay Cave |url=https://www.sciencedirect.com/science/article/pii/S104061822030015X |journal=[[Quaternary International]] |series=Quaternary Stratigraphy and Karst & Cave Sediments: the INQUA-SEQS 2018 Meeting |volume=546 |pages=125–134 |doi=10.1016/j.quaint.2020.01.014 |bibcode=2020QuInt.546..125G |issn=1040-6182 |access-date=22 March 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S104061822030015X |url-status=live }}</ref> and across most of [[northern Eurasia]] into Alaska and [[Yukon]].<ref name="Stuart2011" /> The cave lion had a wide elevation range as well; in Slovakia, [[skull]], [[femur]] and [[pelvis]] remains were excavated in ten [[Karst]] caves in hilly and montane areas at elevations from {{cvt|240|to|1133|m}}.<ref>{{cite journal |author=Sabol, M. |year=2011 |title=A record of Pleistocene lion-like felids in the territory of Slovakia |journal=Quaternaire |series=Hors-série |issue=4 |pages=215−228}}</ref>
-''P. spelaea'' formed a contiguous population from Europe to [[Alaska]] over the [[Bering land bridge]], across the range of the [[mammoth steppe]].<ref>{{cite journal |author=Ersmark, E. |author2=Orlando, L. |author3=Sandoval-Castellanos, E. |author4=Barnes, I. |author5=Barnett, R. |author6=Stuart, A. |author7=Lister, A. |author8=Dalén, L. |year=2015 |title=Population Demography and Genetic Diversity in the Pleistocene Cave Lion |journal=Open Quaternary |volume=1 |issue=1 |pages=Art. 4 |doi=10.5334/oq.aa|doi-access=free }}</ref> It was widely distributed from the [[Iberian Peninsula]], [[Southeast Europe]], [[Great Britain]], [[Central Europe]], the [[East European Plain]], and across most of [[northern Eurasia]] into [[Canada]] and Alaska. The oldest known fossils were excavated in northeastern [[Yakutia]] and were [[radiocarbon date]]d at 62,400 years old. The youngest known fossils are dated 11,925 years old and originated near [[Fairbanks, Alaska]].<ref name=Stuart2011/>
+The oldest known fossils dated with [[radiocarbon dating]] were excavated in northeastern Russia and were radiocarbon dated at 62,400 years old.<ref name="Stuart2011" /> [[Phalanx bone]]s excavated in Spain's [[La Garma cave complex]] were radiocarbon dated to 14,300–14,000 years old.<ref name="UnderTheSkinOfLion" /> The youngest known fossils are dated 11,925 years old and originated near [[Fairbanks, Alaska]].<ref name="Stuart2011" /> In Western Europe, however, ''P. spelaea'' may have survived into the Early Holocene.<ref>{{Cite journal |last1=Masseti |first1=Marco |last2=Mazza |first2=Paul P. A. |date=9 April 2013 |title=Western European Quaternary lions: new working hypotheses: European Quaternary Lions |url=https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/bij.12032 |journal=[[Biological Journal of the Linnean Society]] |language=en |volume=109 |issue=1 |pages=66–77 |doi=10.1111/bij.12032 |access-date=24 March 2024 |archive-date=24 March 2024 |archive-url=https://web.archive.org/web/20240324173452/https://academic.oup.com/biolinnean/article-lookup/doi/10.1111/bij.12032 |url-status=live }}</ref>
-[[Phalanx bone]]s excavated in Spain's [[La Garma cave complex]] were radiocarbon dated to 14,300–14,000 years old.<ref>{{cite journal |author=Cueto, M. |author2=Camarós, E. |author3=Castaños, P. |author4=Ontañón, R. |author5=Arias, P. |year=2016 |title=Under the Skin of a Lion: Unique Evidence of Upper Paleolithic Exploitation and Use of Cave Lion (''Panthera spelaea'') from the Lower Gallery of La Garma (Spain) |journal=PLOS ONE |volume=11 |issue=10 |pages=e0163591 |doi=10.1371/journal.pone.0163591|pmid=27783697 |pmc=5082676 |bibcode=2016PLoSO..1163591C |doi-access=free }}</ref>
-In Slovakia, [[skull]], [[femur]] and [[pelvis]] remains were excavated in ten [[Karst]] caves in hilly and montane areas at elevations from {{convert|240|to|1133|m|ft|abbr=on}}.<ref>{{cite journal |author=Sabol, M. |year=2011 |title=A record of Pleistocene lion-like felids in the territory of Slovakia |journal=Quaternaire |series=Hors-série |issue=4 |pages=215−228 }}</ref>
-In [[Yakutia]]'s Khayrgas Cave, bones of ''P. spelaea'' were found together with remains of humans, wolf, [[reindeer]], Pleistocene [[Equus ferus|horse]] and fish in a layer dated 13,200–21,500 years old.<ref>{{cite journal |author=Kuzmin, Y. V. |author2=Kosintsev, P. A. |author3=Stepanov, A. D. |author4=Boeskorov, G. G. |author5=Cruz, R. J. |year=2017 |title=Chronology and faunal remains of the Khayrgas Cave (Eastern Siberia, Russia) |journal=Radiocarbon |volume=59 |issue= 2|pages=575−582 |doi=10.1017/RDC.2016.39 |bibcode=2017Radcb..59..575K |s2cid=133453976 |url=https://pdfs.semanticscholar.org/a35f/407f5151a0f55c7e1501b7ac4d2966624565.pdf |archive-url=https://web.archive.org/web/20200220004550/https://pdfs.semanticscholar.org/a35f/407f5151a0f55c7e1501b7ac4d2966624565.pdf  |archive-date=2020-02-20 }}</ref>
+In 2008, a well-preserved mature cave lion specimen was unearthed near the [[Maly Anyuy River]] in [[Chukotka Autonomous Okrug]], which still retained some clumps of hair.<ref>{{Cite journal |author=Kirillova, I. V. |author2=Tiunov, A. V. |author3=Levchenko, V. A. |author4=Chernova, O. F. |author5=Yudin, V. G. |author6=Bertuch, F. |author7=Shidlovskiy, F. K. |year=2015 |title=On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia) |journal=[[Quaternary Science Reviews]] |volume=117 |pages=135–151 |bibcode=2015QSRv..117..135K |doi=10.1016/j.quascirev.2015.03.029}}</ref> In the [[Sakha Republic]]'s Khayrgas Cave, bones of ''P. spelaea'' were found together with remains of humans, wolf, [[reindeer]], Pleistocene [[Equus ferus|horse]] and fish in a layer dated 13,200–21,500 years old.<ref>{{cite journal |author=Kuzmin, Y. V. |author2=Kosintsev, P. A. |author3=Stepanov, A. D. |author4=Boeskorov, G. G. |author5=Cruz, R. J. |year=2017 |title=Chronology and faunal remains of the Khayrgas Cave (Eastern Siberia, Russia) |url=https://pdfs.semanticscholar.org/a35f/407f5151a0f55c7e1501b7ac4d2966624565.pdf |journal=[[Radiocarbon (journal)|Radiocarbon]] |volume=59 |issue=2 |pages=575−582 |bibcode=2017Radcb..59..575K |doi=10.1017/RDC.2016.39 |s2cid=133453976 |archive-url=https://web.archive.org/web/20200220004550/https://pdfs.semanticscholar.org/a35f/407f5151a0f55c7e1501b7ac4d2966624565.pdf |archive-date=20 February 2020}}</ref>
+The cave lion probably inhabited predominantly open habitats such as [[steppe]] and grasslands although it would have also have occurred in open woodlands as well.<ref name=Stuart2011/> It also lived in mountainous regions, likely because it sought out hibernating bears in montane caves as a food source during the winter.<ref>{{Cite journal |last1=Sabol |first1=M. |last2=Gullár |first2=J. |last3=Horvát |first3=J. |date=2018 |title=Montane record of the late Pleistocene ''Panthera spelaea'' (Goldfuss, 1810) from the Západné Tatry Mountains (northern Slovakia) |journal=[[Journal of Vertebrate Paleontology]] |volume=38 |issue=3 |pages=e1467921 |doi=10.1080/02724634.2018.1467921 |doi-access=free|bibcode=2018JVPal..38E7921S }}</ref> While during the Last Glacial Period it was often associated with cold environments, the species also inhabited temperate environments,<ref>{{Cite journal |last=Álvarez-Lao |first=Diego J. |last2=Álvarez-Vena |first2=Adrián |last3=Ballesteros |first3=Daniel |last4=García |first4=Nuria |last5=Laplana |first5=César |date=February 2020 |title=A cave lion (Panthera spelaea) skeleton from Torca del León (NW Iberia): Micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3) |url=https://linkinghub.elsevier.com/retrieve/pii/S0277379119306420 |journal=Quaternary Science Reviews |language=en |volume=229 |pages=106123 |doi=10.1016/j.quascirev.2019.106123|hdl=10651/54312 |hdl-access=free }}</ref> such as in Europe during the [[Last Interglacial|Last Interglacial/Eemian]].<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=June 2011 |title=A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord, central Germany |url=http://www.tandfonline.com/doi/abs/10.1080/08912963.2010.507814 |journal=Historical Biology |language=en |volume=23 |issue=2-3 |pages=195–217 |doi=10.1080/08912963.2010.507814 |issn=0891-2963}}</ref>
-In 2008, a well-preserved mature cave lion specimen was unearthed near the [[Maly Anyuy River]] in [[Chukotka Autonomous Okrug]] in [[Russia]], which still retained some clumps of hair.<ref>{{Cite journal |author=Kirillova, I. V. |author2=Tiunov, A. V. |author3=Levchenko, V. A. |author4=Chernova, O. F. |author5=Yudin, V. G. |author6=Bertuch, F. |author7=Shidlovskiy, F. K. |year=2015 |title=On the discovery of a cave lion from the Malyi Anyui River (Chukotka, Russia) |journal=Quaternary Science Reviews |volume=117 |pages=135–151 |doi=10.1016/j.quascirev.2015.03.029|bibcode=2015QSRv..117..135K }}</ref>
-The cave lion was probably predominantly found in open habitats such as [[steppe]] and grasslands although it would have also have occurred in open woodlands as well.<ref name="Stuart2011"/>
 ===Discoveries===
 [[File:Sparta body lateral view.png|thumb|"Sparta", a 28,000 year old mummified female cave lion cub from the banks of the Semyuelyakh River in Siberia.<ref name="Boeskorov_al.2021" />]]
-In 2015, two frozen cave lion cubs, estimated to be between 25,000 and 55,000 years old, were discovered close to the [[Uyandina River]] in Yakutia, [[Siberia]] in [[permafrost]].<ref>{{cite news |url=http://siberiantimes.com/science/casestudy/news/n0796-extinct-lion-cubs-found-in-siberia-are-up-to-55000-years-old-latest-test-results-reveal/|title=Extinct lion cubs found in Siberia are up to 55,000 years old – latest test results reveal |work=The Siberian Times |date=3 November 2016 |author=<!-- Staff -->|access-date=12 November 2017}}</ref><ref>{{cite news |title=Meet this extinct cave lion, at least 10,000 years old – world exclusive | first = Anastasia | last = Koryakina | date = 26 October 2015 |url=http://siberiantimes.com/science/others/news/n0464-meet-this-extinct-cave-lion-at-least-10000-years-old/ |work=The Siberian Times |access-date=29 October 2015}}</ref><ref>{{cite news |title=Frozen Cave Lion Cubs from the Ice Age Found in Siberia |url=https://www.nationalgeographic.com/science/article/151028-cave-lion-frozen-permafrost-science/|work=National Geographic News|publisher=National Geographic|access-date=29 October 2015|first=Riley |last=Black |date=28 October 2015}}</ref>
+In 2015, two frozen cave lion cubs, estimated to be between 25,000 and 55,000 years old, were discovered close to the [[Uyandina River]] in Yakutia, [[Siberia]] in [[permafrost]].<ref>{{cite news|url=http://siberiantimes.com/science/casestudy/news/n0796-extinct-lion-cubs-found-in-siberia-are-up-to-55000-years-old-latest-test-results-reveal/|title=Extinct lion cubs found in Siberia are up to 55,000 years old – latest test results reveal|work=The Siberian Times|date=3 November 2016|author=<!-- Staff -->|access-date=12 November 2017|archive-date=12 November 2017|archive-url=https://web.archive.org/web/20171112190748/http://siberiantimes.com/science/casestudy/news/n0796-extinct-lion-cubs-found-in-siberia-are-up-to-55000-years-old-latest-test-results-reveal/|url-status=live}}</ref><ref>{{cite news |title=Meet this extinct cave lion, at least 10,000 years old – world exclusive |first=A. |last=Koryakina |date=2015 |url=http://siberiantimes.com/science/others/news/n0464-meet-this-extinct-cave-lion-at-least-10000-years-old/ |work=The Siberian Times |access-date=29 October 2015 |archive-date=17 April 2019 |archive-url=https://web.archive.org/web/20190417074356/http://siberiantimes.com/science/others/news/n0464-meet-this-extinct-cave-lion-at-least-10000-years-old/ |url-status=live }}</ref><ref>{{cite news |title=Frozen Cave Lion Cubs from the Ice Age Found in Siberia |url=https://www.nationalgeographic.com/science/article/151028-cave-lion-frozen-permafrost-science/|archive-url=https://web.archive.org/web/20210226152104/https://www.nationalgeographic.com/science/article/151028-cave-lion-frozen-permafrost-science |url-status=dead |archive-date=26 February 2021|work=National Geographic News|publisher=National Geographic |access-date=29 October 2015 |first=R. |last=Black |date=2015}}</ref>
-Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.<ref>{{cite news|url=http://siberiantimes.com/science/casestudy/news/n0491-whiskers-still-bristling-after-more-than-10000-years-in-the-siberian-cold/|title=Whiskers still bristling after more than 12,000 years in the Siberian cold|work=The Siberian Times|date=17 November 2015|author=<!-- Staff -->|access-date=25 November 2016}}</ref>
+Research results indicate that the cubs were likely barely a week old at the time of their deaths, as their milk teeth had not fully erupted. Further evidence suggests the cubs were hidden at a den site until they were strong enough to follow their mother back to the pride, as with modern lions. Researchers believe that the cubs were trapped and killed by a landslide, and that the absence of oxygen underground hindered their decomposition and allowed the cubs to be preserved in such good condition. A second expedition to the site where the cubs were found was planned for 2016, in hopes of finding either the remains of a third cub or possibly the cubs' mother.<ref>{{cite news |url=http://siberiantimes.com/science/casestudy/news/n0491-whiskers-still-bristling-after-more-than-10000-years-in-the-siberian-cold |title=Whiskers still bristling after more than 12,000 years in the Siberian cold |work=The Siberian Times |date=2015 |author=<!-- Staff --> |access-date=25 November 2016 |archive-date=16 December 2016 |archive-url=https://web.archive.org/web/20161216074651/http://siberiantimes.com/science/casestudy/news/n0491-whiskers-still-bristling-after-more-than-10000-years-in-the-siberian-cold/ |url-status=live }}</ref>
-In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River ({{lang-ru|Тирехтях}}), a tributary of the [[Indigirka]] River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.<ref>{{cite news|url=http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/|title=Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes |work=The Siberian Times |date=9 November 2017|author=<!-- Staff -->|access-date=12 November 2017}}</ref> In 2018, another preserved carcass of a cub was found in a location {{cvt|15|m|ft|sigfig=1}} away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.<ref>{{cite news |last=Gertcyk |first=Olga |title=Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost |publisher=The Siberian Times |url=https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |date=12 September 2018 |access-date=16 September 2018}}</ref> However, [[carbon date|carbon dating]] showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.<ref name="Boeskorov_al.2021" /> Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the [[Ice age]].<ref>{{cite news |last=Hunt |first=K. |title=Perfectly preserved cave lion cub found frozen in Siberia is 28,000 years old. Even its whiskers are intact |newspaper=[[CNN]] |url=https://edition.cnn.com/2021/08/05/world/frozen-cave-lion-cubs-siberia-scn/index.html |date=5 August 2021 |access-date=13 August 2021}}</ref><ref>{{cite news |last=Fitzsimons |first=T. |title=Scientists unveil extinct Ice Age lion cubs pulled from Russian permafrost: The scientists believe that the cave lion cubs, dubbed Boris and Sparta, each briefly roamed the steppe of what is now eastern Russia thousands of years ago |newspaper=[[NBC]] News |url=https://www.nbcnews.com/science/science-news/scientists-pull-extinct-cave-lion-cubs-russian-permafrost-n1276137 |date=6 August 2021 |access-date=13 August 2021}}</ref><ref>{{cite news |last=Gertcyk |first=O. |title=Meet Sparta, the 'best preserved ice age animal ever found' |newspaper=[[The Siberian Times]] |url=https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |date=9 August 2021 |access-date=13 August 2021}}</ref><ref>{{cite magazine |last=Kindy |first=D. |title=Near-Perfect Cave Lion Cub Corpse Found in Siberian Permafrost: Scientists discover two mummified cubs estimated to be approximately 28,000 and 43,000 years old |magazine=[[Smithsonian (magazine)|Smithsonian Magazine]] | publisher=[[Smithsonian Institution]] |url=https://www.smithsonianmag.com/smart-news/near-perfect-cave-lion-cub-corpse-found-siberian-permafrost-180978403/ |date=10 August 2021 |access-date=13 August 2021}}</ref>
+In 2017, another frozen specimen, thought to be a lion cub, was found in Yakutia on the banks of the Tirekhtyakh River ({{lang-ru|Тирехтях}}), a tributary of the [[Indigirka]] River. This male cub was thought to be slightly older than the 2015 cubs at the time of its death; it is estimated to have been around one and a half to two months.<ref>{{cite news |url=http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |title=Extinct cave lion cub in 'perfect' condition found in Siberia rising cloning hopes |work=The Siberian Times |date=2017 |author=<!-- Staff --> |access-date=12 November 2017 |archive-date=12 November 2017 |archive-url=https://web.archive.org/web/20171112012143/http://siberiantimes.com/science/casestudy/news/extinct-cave-lion-cub-in-perfect-condition-found-in-siberia-rising-cloning-hopes/ |url-status=live }}</ref> In 2018, another preserved carcass of a cub was found in a location {{cvt|15|m|ft|sigfig=1}} away. It was considered to be around a month old when it died approximately 50,000 years ago, and presumed to be a sibling of the male cub.<ref>{{cite news |last=Gertcyk |first=O. |title=Cute first pictures of new 50,000 year old cave lion cub found perfectly preserved in permafrost |publisher=The Siberian Times |url=https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |date=2018 |access-date=16 September 2018 |archive-date=14 September 2018 |archive-url=https://web.archive.org/web/20180914132515/https://siberiantimes.com/science/casestudy/news/cute-first-pictures-of-new-50000-year-old-cave-lion-cub-found-perfectly-preserved-in-permafrost-of-yakutia/ |url-status=live }}</ref> However, [[carbon date|carbon dating]] showed them to have lived about 15,000 years apart, with the female estimated to have lived 28,000 years ago, and the male 43,448 years ago.<ref name="Boeskorov_al.2021" /> Both cubs were well preserved, albeit with a few damages, with the female possibly being the "best preserved" animal discovered from the [[Ice age]].<ref>{{cite news |last=Gertcyk |first=O. |title=Meet Sparta, the 'best preserved ice age animal ever found' |newspaper=[[The Siberian Times]] |url=https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |date=9 August 2021 |access-date=13 August 2021 |archive-date=12 August 2021 |archive-url=https://web.archive.org/web/20210812082545/https://siberiantimes.com/other/others/news/meet-sparta-the-best-preserved-ice-age-animal-ever-found/ |url-status=live }}</ref>
 ==Paleobiology==
 [[File:Hoehlenloewe CaveLion hharder.jpg|thumb|right|A cave lion with a reindeer, painting by [[Heinrich Harder]]<ref>{{cite book |author=Bölsche, W. |name-list-style=amp |author2=Harder, H. |year=1900 |title=Tiere der Urwelt. Serie III |location=Wandsbek-Hamburg |publisher=Verlag der Kakao-Compagnie Theodor Reichardt}}</ref>]]
-''P. spelaea'' inhabited open environments such as [[mammoth steppe]] and [[boreal forest]]. It was one of the [[keystone species]] of the mammoth steppe, being one of the main apex predators alongside [[gray wolf]], [[cave hyena]] and [[brown bear]].<ref name=Bocherens2015>{{Cite journal|last=Bocherens|first=H. |date=2015 |title=Isotopic tracking of large carnivore palaeoecology in the mammoth steppe |journal=Quaternary Science Reviews |volume=117 |pages=42–71 |doi=10.1016/j.quascirev.2015.03.018 |bibcode=2015QSRv..117...42B |url=https://www.researchgate.net/publication/275219303}}</ref> Large amounts of bones belonging to ''P. spelaea'' were excavated in caves, where bones of cave hyena, [[cave bear]] and [[Paleolithic]] artefacts were also found.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=The largest European lion ''Panthera leo spelaea'' (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe |journal=Historical Biology |volume=23 |issue=2–3 |pages=271–311 |doi=10.1080/08912963.2010.546529 |s2cid=86638786}}</ref><ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Pleistocene ''Panthera leo spelaea'' (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites |journal=Quaternaire |volume=22 |issue=2 |pages=105–127 |doi=10.4000/quaternaire.5897 |url=http://journals.openedition.org/quaternaire/5897}}</ref>
+''P. spelaea'' inhabited colder environments such as [[mammoth steppe]] and [[boreal forest]]. It was one of the [[keystone species]] of the mammoth steppe, being one of the main [[apex predator]]s alongside the [[gray wolf]], [[cave hyena]] and [[brown bear]].<ref name=Bocherens2015>{{Cite journal |last=Bocherens |first=H. |date=2015 |title=Isotopic tracking of large carnivore palaeoecology in the mammoth steppe |journal=[[Quaternary Science Reviews]] |volume=117 |pages=42–71 |doi=10.1016/j.quascirev.2015.03.018 |bibcode=2015QSRv..117...42B |url=https://www.researchgate.net/publication/275219303 |access-date=3 September 2020 |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518072012/https://www.researchgate.net/publication/275219303_Isotopic_tracking_of_large_carnivore_palaeoecology_in_the_mammoth_steppe |url-status=live }}</ref> Large amounts of bones belonging to ''P. spelaea'' were excavated in caves, where bones of cave hyena, [[cave bear]] and [[Paleolithic]] artefacts were also found.<ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=The largest European lion ''Panthera leo spelaea'' (Goldfuss 1810) population from the Zoolithen Cave, Germany: specialised cave bear predators of Europe |journal=Historical Biology |volume=23 |issue=2–3 |pages=271–311 |doi=10.1080/08912963.2010.546529 |bibcode=2011HBio...23..271D |s2cid=86638786}}</ref><ref>{{cite journal |author=Diedrich, C. G. |year=2011 |title=Pleistocene ''Panthera leo spelaea'' (Goldfuss 1810) remains from the Balve cave (NW Germany) - a cave bear, hyena den and middle palaeolithic human cave – and review of the Sauerland Karst lion cave sites |journal=Quaternaire |volume=22 |issue=2 |pages=105–127 |doi=10.4000/quaternaire.5897 |url=http://journals.openedition.org/quaternaire/5897 |access-date=18 September 2020 |archive-date=11 April 2021 |archive-url=https://web.archive.org/web/20210411153241/https://journals.openedition.org/quaternaire/5897 |url-status=live }}</ref> Some of these accumulations of cave lion bones have been attributed to hoarding of meat from cave lion carcasses by cave hyenas in caves occupied by the latter.<ref>{{Cite journal |last=Diedrich |first=Cajus G. |date=1 July 2011 |title=Late Pleistocene steppe lion Panthera leo spelaea (Goldfuss, 1810) footprints and bone records from open air sites in northern Germany – Evidence of hyena-lion antagonism and scavenging in Europe |url=https://www.sciencedirect.com/science/article/pii/S0277379111000783 |journal=[[Quaternary Science Reviews]] |volume=30 |issue=15 |pages=1883–1906 |doi=10.1016/j.quascirev.2011.03.006 |bibcode=2011QSRv...30.1883D |issn=0277-3791 |access-date=4 February 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071940/https://www.sciencedirect.com/science/article/abs/pii/S0277379111000783 |url-status=live }}</ref> It is unclear whether ''P. spelaea'' was social like the modern lion; some evidence indicates that it may have been solitary.<ref name=Bocherens2015/>
-It is unclear whether ''P. spelaea'' was social like the modern lion; some evidence indicates that it may have been solitary.<ref name=Bocherens2015/>
+Isotopic analyses of bone [[collagen]] samples extracted from fossils indicate that cave bear cubs, [[reindeer]] and other [[cervid]]s (deer) were prominent in the diet of cave lions. Later cave lions seem to have preyed foremost on reindeer, up to the brink of local extinction or extirpation of both species.<ref name="Bocherens2015" /><ref name=Bocherens>{{cite journal |author1=Bocherens, H. |name-list-style=amp |author2=Drucker, D. G. |author3=Bonjean, D. |author4=Bridault, A. |author5=Conard, N. J. |author6=Cupillard, C. |author7=Germonpré, M. |author8=Höneisen, M. |author9=Münzel, S. C. |author10=Napierala, H. |author11=Patou-Mathis, M. |date=2011 |title=Isotopic evidence for dietary ecology of cave lion (''Panthera spelaea'') in North-Western Europe: prey choice, competition and implications for extinction |journal=Quaternary International |volume=245 |issue=2 |pages=249–261 |url=https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |doi=10.1016/j.quaint.2011.02.023 |bibcode=2011QuInt.245..249B |s2cid=129706533 |access-date=18 September 2020 |archive-date=11 May 2020 |archive-url=https://web.archive.org/web/20200511073456/https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |url-status=live }}</ref> Other possible prey species were [[giant deer]], [[red deer]], [[wild horse]], [[muskox]], [[aurochs]], [[wisent]], [[bison priscus|steppe bison]], young [[woolly rhino]], and young [[woolly mammoth]]. It likely competed for prey with the [[Panthera pardus spelaea|European Ice Age leopard]], cave hyenas, brown bears and grey wolves in Eurasia,<ref name="Diedrich 2013">{{cite journal |author=Diedrich, C. G. |year=2013 |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |pages=167–193 |doi=10.1016/j.quascirev.2013.05.009 |volume=76|bibcode=2013QSRv...76..167D}}</ref> along with [[Arctodus|short-faced bears]], ''[[Homotherium]]'', and [[Beringian wolf|Beringian wolves]] in [[Beringia]].<ref name="Bocherens2015" />
+== Extinction ==
-Isotopic analyses of bone [[collagen]] samples extracted from fossils indicate that cave bear cubs, [[reindeer]] and other [[cervid]]s were prominent in the diet of cave lions. Later cave lions seem to have preyed foremost on reindeer, up to the brink of local extinction or extirpation of both species.<ref name=Bocherens>{{cite journal |author1=Bocherens, H. |name-list-style=amp |author2=Drucker, D. G. |author3=Bonjean, D. |author4=Bridault, A. |author5=Conard, N. J. |author6=Cupillard, C. |author7=Germonpré, M. |author8=Höneisen, M. |author9=Münzel, S. C. |author10=Napierala, H. |author11=Patou-Mathis, M. |date=2011 |title=Isotopic evidence for dietary ecology of cave lion (''Panthera spelaea'') in North-Western Europe: prey choice, competition and implications for extinction |journal=Quaternary International |volume=245 |issue=2 |pages=249–261 |url=https://hal.archives-ouvertes.fr/hal-01673488/file/BOCHERENS%20%26%20al%202011%28QI-lion%29.pdf |doi=10.1016/j.quaint.2011.02.023 |bibcode=2011QuInt.245..249B |s2cid=129706533}}</ref> Other possible prey species were [[giant deer]], [[red deer]], [[wild horse]], [[muskox]], [[aurochs]], [[wisent]], [[bison priscus|steppe bison]], young [[woolly rhino]] and young [[woolly mammoth]]. It likely competed for prey with the European Ice Age leopard ([[Panthera pardus spelaea|''P. pardus spelaea'']]) as well as cave hyenas, cave bears, gray wolves and in North America, [[Arctodus|short-faced bears]] and [[dire wolf|dire wolves]].<ref name="Diedrich 2013">{{cite journal |author=Diedrich, C. G. |year=2013 |title=Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art |journal=Quaternary Science Reviews |pages=167–193 |doi=10.1016/j.quascirev.2013.05.009 |volume=76|bibcode=2013QSRv...76..167D}}</ref> An Isotope analysis study suggested most sampled ''P. spelea'' specimens were primarily consuming [[reindeer]].<ref name=Bocherens2015 />
+The extinction of the Eurasian cave lion was multifaceted, being attributable to a combination of human activity, the decline of the mammoth steppe, and changes in the herbivore guild.<ref name=":0">{{Cite journal |last1=Marciszak |first1=Adrian |last2=Lipecki |first2=Grzegorz |last3=Pawłowska |first3=Kamilla |last4=Jakubowski |first4=Gwidon |last5=Ratajczak-Skrzatek |first5=Urszula |last6=Zarzecka-Szubińska |first6=Katarzyna |last7=Nadachowski |first7=Adam |date=20 December 2021 |title=The Pleistocene lion Panthera spelaea (Goldfuss, 1810) from Poland – A review |url=https://www.sciencedirect.com/science/article/pii/S1040618220308399 |journal=[[Quaternary International]] |series=The Quaternary of Europe and Adjacent Areas: Stratigraphical Perspectives and Tools for Correlations – SEQS-2019 |volume=605-606 |pages=213–240 |doi=10.1016/j.quaint.2020.12.018 |bibcode=2021QuInt.605..213M |issn=1040-6182 |access-date=22 March 2024 |via=Elsevier Science Direct |archive-date=18 May 2024 |archive-url=https://web.archive.org/web/20240518071941/https://www.sciencedirect.com/science/article/abs/pii/S1040618220308399 |url-status=live }}</ref> There is direct evidence of human hunting of cave lions and exploitation of their pelts for ritualistic purposes during the [[Magdalenian]], which was likely an important factor contributing to their ultimate demise.<ref name="UnderTheSkinOfLion">{{Cite journal |last1=Cueto |first1=Marián |last2=Camarós |first2=Edgard |last3=Castaños |first3=Pedro |last4=Ontañón |first4=Roberto |last5=Arias |first5=Pablo |date=26 October 2016 |editor-last=Macchiarelli |editor-first=Roberto |title=Under the Skin of a Lion: Unique Evidence of Upper Paleolithic Exploitation and Use of Cave Lion (Panthera spelaea) from the Lower Gallery of La Garma (Spain) |journal=[[PLOS ONE]] |language=en |volume=11 |issue=10 |pages=e0163591 |doi=10.1371/journal.pone.0163591 |doi-access=free |issn=1932-6203 |pmc=5082676 |pmid=27783697|bibcode=2016PLoSO..1163591C }}</ref>
 ==See also==
 {{div col|colwidth=15em}}
 *[[History of lions in Europe]]
+*[[Megafauna]]
 *''[[Panthera atrox]]''
 *''[[Panthera blytheae]]''